Download A preliminary biological survey of Cerro Piedra Larga, Oaxaca, Mexico

Anales del Instituto de Biología, Universidad Nacional Autónoma de México,
Serie Zoología 75(2): 439-466. 2004
A preliminary biological survey of Cerro Piedra Larga,
Oaxaca, Mexico: Birds, mammals, reptiles, amphibians, and
plants
A. TOWNSEND PETERSON*
LUIS CANSECO MÁRQUEZ**
JOSÉ LUIS CONTRERAS JIMÉNEZ***
GRISELDA ESCALONA-SEGURA****
OSCAR FLORES-VILLELA**
JOSÉ GARCÍA-LÓPEZ*****
BLANCA HERNÁNDEZ-BAÑOS**
CARLOS A. JIMÉNEZ RUIZ**
LIVIA LEÓN-PANIAGUA**
SERGIO MENDOZA AMARO**
ADOLFO G. NAVARRO-SIGÜENZA**
VICTOR SÁNCHEZ-CORDERO*****
DAVID E. WILLARD******
Resumen. Cerro Piedra Larga es una montaña aislada en el este de Oaxaca,
justo al oeste del Istmo de Tehuantepec. En virtud de que en esta sierra no se
ha conducido ningún inventario biológico, un estudio preliminar puede ser de
interés. Durante varias semanas de trabajo en las partes altas de la sierra,
desarrollamos inventarios preliminares para aves, mamíferos, reptiles, anfibios
y plantas. Geográficamente, las afinidades de la biota de la sierra parecen
constituir una mezcla entre la Sierra Madre Oriental y la Sierra Madre del Sur,
lo cual sugiere una origen por medio de colonización y no por medio de
conexiones históricas.
Palabras clave: inventario biológico, faunística, florística, Cerro Piedra Larga,
Oaxaca, México.
* Natural History Museum, The University of Kansas, Lawrence, Kansas 66045.
** Facultad de Ciencias, Universidad Nacional Autónoma de Mexico, Apartado postal 70-399, 04510
México, D.F., México.
*** Herbario de la BUAP, Edif. 76 Unidad de Ciencias C.U., 72590 Puebla, Pue., México.
**** El Colegio de la Frontera Sur, Calle 10 #264 por Calle 61, Col. Centro, 24000 Campeche,
Campeche, México.
***** Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México,
Apartado postal 70-153, 04510 México, D. F. México.
****** Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois,
60605, USA.
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A. TOWNSEND PETERSON ET AL.
Abstract. Cerro Piedra Larga is an isolated mountain massif in eastern Oaxaca,
lying just west of the Isthmus of Tehuantepec. No previous biological inventories had assessed this range, making even a preliminary assessment of interest.
During several weeks of work in the higher portions of the mountain range, we
assembled such preliminary inventories for birds, mammals, reptiles, amphibians, and plants. Geographically, the affinities of the highland biota of Cerro
Piedra Larga appear to be mixed between the Sierra Madre Oriental and the
Sierra Madre del Sur, suggesting that the biota likely originated by colonization,
rather than via historical connections.
Key words: biological inventory, faunistics, floristics, Cerro Piedra Larga, Oaxaca,
México.
Introduction
Mexico, located at the conjunction of two great biogeographic realms, is considered
a megadiverse country, one of the storehouses of biodiversity at a global scale
(Mittermeier et al. 1998). Although Mexico has seen attention from students of
biodiversity over more than two centuries (Ramamoorthy et al. 1993), the survey of
its biodiversity remains incomplete (Peterson et al. 1998). Many areas remain
unsurveyed, and incompletely known as to species composition and communities
of animals and plants.
Cerro Piedra Larga is an isolated mountain massif in eastern Oaxaca (Figs. 1,
2), lying less than 100 km west of the Isthmus of Tehuantepec, a major biogeographic
barrier that divides the heart of Mexico from northern Central America. The
montane habitats of the Cerro are isolated from the Sierra de los Mixes and the
Zempoaltepec massif to the north by the dry lowlands of the Río Tehuantepec and
from the Sierra de Miahuatlán to the south and west by a low, dry valley that holds
the highway from Oaxaca City to the Isthmus. Hence, the Cerro represents an
island of montane habitat separated from larger sierras by dry lowland valleys,
and at the extreme of a major biogeographic realm. In spite of its intriguing
geographic situation, Cerro Piedra Larga has seen no organized biological survey,
and in fact has seen effectively no collecting whatsoever in the high montane
portions (Binford 1989).
Initial surveys of Cerro Piedra Larga by small plane revealed a surprise in its
vegetation. Whereas most interior mountain massifs in Mexico simply hold arid
and humid pine-oak forest, Cerro Piedra Larga appeared to hold patches of cloud
forest in high-elevation sheltered canyons. This observation motivated on-theground fieldwork, and was indeed confirmed by our field studies. This patch of
cloud forest is clearly one of the most isolated in all of Mexico, and for that reason
is of considerable biological interest.
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441
Fig. 1. Map of southern Mexico, showing elevational variation (white-to-black scale indicates
elevations in 500 m intervals) and the location of Cerro Piedra Larga.Fig. 3. Species
Cerro Piedra Larga is a mass of Mesozoic origin located just west of the Isthmus of Tehuantepec, northwest of Juchitán, in the area bordered by the coordinates 16o 31’ and 16o 37’ N latitude, and 95o 45’ and 95o 51’ W longitude. The
massif rises from a base at about 600 m to elevations of about 2700 m, and has a
roughly oblong shape.
The present contribution is intended as a multi-taxon approach to a first-pass
documentation of the biological diversity of Cerro Piedra Larga. A team of
investigators and students from the Universidad Nacional Autónoma de México
and the Field Museum of Natural History surveyed the high montane portions of
the Cerro in March, April, and August of 1993. Herein, we present preliminary
inventories for plants, reptiles and amphibians, birds, and mammals.
Inventories for each taxon are clearly preliminary. Species accumulation curves
for the three vertebrate taxa (Fig. 3) indicate that inventories for birds are likely
complete. Mammals, although presenting a flat accumulation curve, were sampled
in a small area, and the true fauna is doubtless larger. Reptiles and amphibians,
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which present a steeply ascending species accumulation curve, are only partly
sampled, and will certainly reveal many additional forms with further study.
Methods
Fig. 2. Map of the region of Cerro Piedra Larga, showing principal access roads, and study
sites (as large black dots).
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
443
Fig. 3. Species accumulation curves showing the progress of each inventory (birds, mammals,
and reptiles and amphibians).
Diverse methods were employed to assemble inventories of species for plants, reptiles and amphibians, birds and mammals. As such, the information (or the depth
of information) available for the different taxonomic groups varies in completeness and detail. In general, our studies were carried out by multiple investigators
per taxon, during 15-day periods in 1993.
Plants. Field work and collections were made on Cerro Piedra Larga in April and
August 1993, each visit lasting 15 days by three botanists. We made systematic
sweeps through the areas surrounding the study sites, focusing on the best-preserved
vegetation patches, and collecting specimens of plants that were either flowering
or fruiting. Appropriate data on date, locality, and habitat were noted to permit
proper documentation and identification in the herbarium.
We developed a vegetation profile based on a representative area with a wellpreserved pine forest. We delimited an area of 5 x 50 m, and censused all trees with
DBH of ˆ3.18 cm: we recorded total height (m), circumference at breast height
(cm), crown coverage (m) via measurement of major and minor perpendicular
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diameters, identification and position within the study area via coordinates in a set
of 5 x 5 m cells.
Reptiles and amphibians. Field work and collections were made on Cerro Piedra
Larga in April and August 1993, each visit lasting 15 days by three herpetologists.
Daily activities included following line transects during approximately 9 hr,
beginning late in the morning, and including the afternoon, evening, and part of
the night. These efforts were designed to include hours of peak activity of most
amphibian and reptile species. In each daily transect, we inspected each microhabitat
represented, attempting each day to include different vegetation types and
altitudinal zones. Two principal vegetation types —pine-oak forest and tropical
deciduous forest (hereafter “dry forest”)— were worked most intensively (Fig. 1).
Specimens were collected for the purpose of vouchering identification and
permitting additional study. Collections were made by traditional capture techniques
(Casas-Andreu et al. 1996), complemented by the use of drift-fences and pitfall
traps (Vogt & Hine 1982). For each specimen collected, we recorded locality, date,
hour of capture, elevation, vegetation type, microhabitat, sex, age, coloration, and
activities observed. Specimens were preserved according to accepted herpetological
techniques (Pisani & Villa 1974), and were deposited in the Colección Herpetológica,
Museo de Zoología “Alfonso L. Herrera,” Facultad de Ciencias, Universidad
Nacional Autónoma de México (MZFC).
Birds. Four experienced observers and two assistants inventoried the birds of Cerro
Piedra Larga from 31 March to 13 April 1993. Two camps were established in
mixed humid pine-oak and cloud forest near Aserradero El Aguacate, near San
Sebastián Jilotepec, Municipio de Nejapa de Madero, Distrito de Yautepec, Oaxaca,
at 2520 m (16° 36.469’ N: 95° 48.045’ W). Geographic coordinates were determined
with a precision of better than 100 m using a Trimble Ensign geographic positioning
system.
Ten to 17 mist nets were set in lines of 4-10, crossing each major microhabitat
type (humid pine-oak forest, cloud forest, edge habitats). Lines were run for 3-7
days, and then shifted in position, for a total of 169 net-days. Additional records
were accumulated by visual observations and by selective hunting. Specimens
resulting from these collections were deposited in the Museo de Zoología, Facultad
de Ciencias, UNAM, the Field Museum, and the Natural History Museum, University
of Kansas.
Mammals. Two experienced mammalogists inventoried the mammals of Cerro
Piedra Larga from 31 March to 13 April 1993. Two camps were established in
mixed humid pine-oak and cloud forest near Aserradero El Aguacate, near San
Sebastián Jilotepec, Municipio de Nejapa de Madero, Distrito de Yautepec, Oaxaca:
(1) Aserradero El Aguacate (16° 36.469’ N: 95° 48.045’ W, 2520 m, pine-oak forest;
7-13 April 1993), and (2) Agua Fría (16° 36.750’ N: 96° 48.750’ W, 2100-2300 m,
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
445
pine-oak forest with isolated patches of cloud forest; 31 March-7 April 1993). Geographic coordinates were determined with a precision of better than 100 m using
a Trimble Ensign geographic positioning system. Work was carried out near the
end of the dry season (< 100 mm per month), so seed and fruit production were
minimal at the time of the study.
Methods employed were the traditional inventory approaches for small-and
medium-sized mammals. Ten 12 m nylon mist nets were opened from 18:00-05:00
hr daily, and were checked every 2 hr, for a total of approximately 5000 net-hr of
sampling in each habitat. Bats captured were kept in cloth bags in shade until they
could be identified and prepared as specimens, and to obtain a sample of their
excretions.
We set 160 Sherman traps baited with oatmeal and vanilla along two transects
of 80 traps each. We also set two transects of pitfall traps, with 11 buckets buried
every 5 m to create a 50 m sampling transect; the buckets were connected via a
barrier of plastic that was buried a few centimeters into the soil to guide shrews
into the pitfall traps. Medium-sized species were captured using Tomahawk traps
baited with sardines. Specimens resulting were deposited in the Museo de Zoología
de la Facultad de Ciencias (MZFCUNAM) y en la Colección Nacional de Mamíferos,
Instituto de Biología (CNM-IBUNAM), both in the Universidad Nacional Autónoma
de México.
Results
Plants. The lower parts of the Cerro are covered with deciduous tropical scrub
(Rzedowski 1978). This community is typical of seasonal climates, and is
characterized by floristic elements that loose foliage during the dry season. It is
dominated by trees that average 5-8 m tall, with some emergents reaching 12 m.
Among dominant families are Burseraceae, Leguminosae, Anacardiaceae, and
Euphorbiaceae. The shrub layer is 0.5-2 m tall, and is quite diverse, including
many species of Leguminosae, Euphorbiaceae, and Verbenaceae. Principal tree
species include Bursera bicolor, B. bipinnata, Haematoxylum brasiletto, Ceiba parvifolia,
Acacia farnesiana, A. cochliacantha, A. pennatula, Alvaradoa amorphoides, Stemmadenia
obovata, Guazuma ulmifolia, Lysiloma divaricata, Euphorbia schlechtendalii, Leucaena
esculenta, Gyrocarpus jatrophifolius, Caesalpinia esclerocarpa, and Bucida wigginsiana.
Between 1800 and 2200 m, a pine-oak forest dominates, with trees 12-25 m
tall, and varying in species composition by aspect and by substrate characteristics.
On southeast-facing rocky slopes, it is common to encounter Pinus lawsonii, P. pringlei,
and P. oocarpa, associated with Quercus crassifolia, Q. acutifolia, Q. scytophylla, Arbutus
xalapensis, Nolina longifolia, Iresine celosia, and others. This zone has seen frequent
forest fires started for creating pastures and facilitating extraction of wood. For
this reason, large open areas in the process of regeneration are common, often
with many individuals of Pinus spp. 5-10 m tall and a few taller adult examples.
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Flowering plants include some orchids, a few Piperaceae, Ericaceae such as Xolisma
squamulosa, Arctostaphylos pyrifolia, Bromeliaceae such as Pitcairnia sp., and others.
At 2200-2600 m, on protected northwestern slopes, and in valleys, a forest
almost purely of Pinus is present, with individuals 15-30 m tall. Among the most
important species are Pinus maximinoi, Pinus douglasiana and P. ayacahuite. In more
open areas, these species intermingle with Quercus scytophylla, Q. candicans, Q. laurina,
Alnus spp., Arbutus xalapensis, A. glandulosa, Styrax argenteus, Myrica cerifera, Rapanea
juergensenii, and Clethra macrophylla. The shrub and herbaceous layers include species
such as Satureja macrostema, Salvia lavanduloides, Salvia mexicana, Litsea glaucescens,
Arctostaphylos pyrifolia, Vaccinium confertum, Xolisma squamulosa, Orthrosanthus
monadelphus, Lotus repens, and Lobelia laxiflora. Fig. 4 shows a diagram of a typical
profile of this sort of forest.
The cloud forest is well-developed only in canyons and on humid northwestfacing slopes, between 2200 and 2600 m. This habitat does not occupy a great area,
and is restricted to disjunct patches of variable floristic composition. It has two arboreal
strata, the taller being made up of 20-30 m trees including Pinus ayacahuite, P.
maximinoi, Quercus uxoris, Q. laurina, and Q. candicans. The lower stratum includes
12-18 m trees, and is dominated by Cornus disciflora, C. excelsa, Garrya laurifolia, Fuchsia
arborescens, Styrax argenteus, Hedyosmum mexicanum, Myrica cerifera, Alnus spp., Budleia
crotonoides, Clethra licanioides, Senecio aspatensis, Inga sp., Cleyera theaeoides, and Saurauia
scabrida. The shrub and herbaceous layers are rich in tropical species, including
Litsea glaucescens, Salvia gracilis, Eugenia oerstediana, Solanum pubigerum, Psychotria sp.,
Palicourea padifolia, and Lycianthes sp. Also common are vines such as Solandra sp.,
Passiflora sexflora, Passiflora membranacea, Smilax regelii, Mikania pyramidata, Marsdenia
macrophylla, and Sarcostemma sp. Epiphytes are another important element, including members of the families Bromeliaceae (e.g., Tillandsia spp.), Orchidaceae (e.g.,
Lemboglossum cervantesii), and Piperaceae (e.g., Peperomia sp.).
Reptiles and amphibians. In all, 34 species were encountered on Cerro Piedra Larga,
including eight amphibians of five genera and four families, and 26 reptiles of 21
genera and 12 families (Table 1, Appendix 2). Of species collected, five were taxa
endemic to the state of Oaxaca: Abronia oaxacae, Phyllodactylus muralis, Tantilla striata,
Micrurus ephippifer, and Ctenosaura oaxacana. Hence, Oaxaca endemics represented
14.7% of the total fauna.
Species detected were fairly evenly divided between the two habitats (Table 2).
Pine-oak forest held 15 species, including snakes (six species) for 40% of the fauna,
lizards (five species) 33.3 %, anurans (three species) 20%, and a salamander (6.6%).
All species detected in this habitat were diurnal, except the crepuscular Rhadinaea
fulvivittis. Several species were tree living, including Abronia oaxacae, Sceloporus
grammicus, and Leptophis diplotropis; Chiropterotriton sp. was found in a bromeliad.
In dry forest, 21 species were found, including lizards (five species) at 23.9% of
the fauna, anurans (six species) 28.5%, and snakes (five species) 23.8%. Of species
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
447
Table 1. Taxonomic diversity of reptiles and amphibians detected in biotic inventories on
Cerro Piedra Larga, Oaxaca.
Taxonomic group
Salamanders
Anurans
Lizards
Snakes
Turtles
Totals
Families
Genera
Species
% of total species
1
4
10
10
1
26
1
7
15
10
1
34
2.9
20.0
42.8
31.4
2.9
100.0
1
3
7
4
1
16
encountered, Phyllodactylus muralis was nocturnal, and Bufo canaliferus was found
both in daytime and nighttime. Trimorphodon biscutatus was found in daytime in
molt, but its activity is usually nocturnal.
Of the 34 species encountered on Cerro Piedra Larga, 27 were restricted to
single habitat types (Table 3). The area representing the ecotone between the two
Table 2. Percentages of representation of taxa of amphibians and reptiles divided by
vegetation type. Percentages given in columns are relative to the column totals, whereas
those given with the column totals refer to the overall total of species encountered.
Taxonomic group
Pine-oak forest
Salamanders
Anurans
Lizards
Snakes
Turtles
Total
1 (6.6%)
3 (20.0%)
5 (33.3%)
6 (40.0%)
.
15 (44.1%)
Dry forest
Ecotone
.
6 (28.5%)
7 (47.6%)
5 (23.8%)
.
21 (61.7%)
.
.
2 (50.0%)
1 (25.0%)
1 (25.0%)
4 (11.7%)
habitats held only four species, of which three were also found in dry forest, and
two in pine-oak forest; only the turtle Rhinoclemmys rubida was detected solely in
the ecotone.
Many species were found on Cerro Piedra Larga only within limited elevational
ranges (Table 3, Fig. 5). For example, the lizards Sceloporus formosus and S. grammicus
were found in pine-oak forest, but only below 2620 m. The lizard Anolis quercorum
was found only at 1850-2095 m. Similarly, in dry forest, the lizard Sceloporus siniferus
was found only at 900-1765 m. Numerous such examples were obtained, in which
a species’ elevational range was less than that of its habitat.
The distribution of species among different microhabitats is of special interest
to understanding the structure of the community in which they live. Reptile and
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amphibian species were found to use four principal microhabitats on Cerro Piedra
Larga, including on and below the forest floor, in trees, and on rock walls (Table 4).
Twenty species inhabited the forest floor, on and under rocks, on or below fallen
logs, in leaf litter, in fenceposts, in grass, or among magueys. Some species, such as
Sceloporus siniferus, were observed to use numerous niches on the forest floor. Among
arboreal species, the salamander Chiropterotriton sp. was found in a bromeliad; other
species were reptiles found in shrubs, in trees, or on fenceposts. Fossorial species
included several snakes, which were found typically beneath rocks and trunks;
finally, the lizard Urosaurus bicarinatus used rock walls extensively, in addition to
the forest floor.
It is important to point out that Cerro Piedra Larga was visited only twice for
herpetological studies, owing in large part to the difficulty of access to the higher
Fig. 4. Vegetation profile of Cerro Piedra Larga. See text for explanation.
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
449
portions of the mountain. The total of 34 species detected thus compares favorably with 26 species in the vicinity of San Cristóbal de las Casas (Hernández M.
1992), and 37 species in Omiltemi, Guerrero (Flores V. & Muñoz 1993). Although
this comparison indicates that the zone is probably herpetologically quite rich, the
preliminary nature of the inventory reported herein must be emphasized.
Three of the species documented herein represent extensions to known
geographic distributions. The snake Tantilla striata was previously known from a
single locality 61 km NE of Cerro Mixtequilla, Oaxaca (Wilson 1990). The record
of the frog Hyla bistincta may represent the southernmost record of the species,
which is otherwise known from localities in the northern highlands of Oaxaca
(Duellman 2001). Finally, the presence of the lizard Abronia oaxacae on Cerro Piedra
Larga represent an eastward extension of the species’ known distribution, the nearest
known localities being Santo Domingo Chontecomatlán, in the Sierra Madre del
Sur, and Sierra de Juárez, in northern Oaxaca (Cambell & Frost 1993).
No plethodontid salamander of the genus Chiropterotriton was indicated for the
Cerro Piedra Larga region in the most recent taxonomic revision (Darda 1994),
probably for lack of material for study from the area. The individual collected on
Cerro Piedra Larga belongs to the “Southern Group” (Darda 1994). Previous Oaxaca
records included the southern extreme of the Sierra Madre Oriental, and a probable
population in the Sierra Madre del Sur. Because the relationships and taxonomic
Fig. 5. Elevational distribution of reptile and amphibian species on Cerro Piedra Larga,
Oaxaca. Numbers refer to species in Appendix 2. Oaxaca endemic species are indicated
with bold numbers. DF = dry forest, POF = pine-oak forest, SSJ = San Sebastián Jilotepec.
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status of these populations remains problematic, the specific identification of the
individual collected on Cerro Piedra Larga must await more detailed study.
Some reptile and amphibian species on Cerro Piedra Larga were found only in
the pine-oak forest, and others principally in the dry forest; one was found only in
the ecotone between the two habitats. These patterns clearly are the product of sets
of causal factors, including both biotic factors (e.g., vegetation characteristics,
competitive and mutualistic interactions with other species) and abiotic factors
(e.g., altitude, humidity, etc.).
An excellent example of the effect of these ecological restrictions is that of the
Chiropterotriton salamanders, which are typically found only in situations of high
humidity in forests with abundant epiphytes. Wake (1987) indicated that most of
these salamanders inhabit bromeliads, probably because of abundant food resources,
egg deposition sites, and buffering against extremes of temperature and humidity.
In contrast, Abronia oaxacae is usually considered restricted to pine-oak forests
(Bogert & Porter 1967). As for dry forest, several species were found that appear
restricted to arid zones, including Urosaurus bicarinatus and Sceloporus siniferus
(Duellman 1966, Sánchez & López-Forment 1988).
Because the pine-oak forest on Cerro Piedra Larga is surrounded by dry forest,
it is effectively an island, probably isolated during the climatic fluctuations related
to glaciations in the Pleistocene. The pine-oak forests in the area are distinct from
the surrounding habitats, suggesting differentiation of the faunas of the habitats.
Hence, we consider the fauna of the pine-oak forests of Cerro Piedra Larga to
represent an isolated montane fauna.
Twelve of the 15 species found in this habitat were not found in the adjacent
dry forest (including an unidentified Hyla tadpole and a shed Senticolis triapis skin).
Of these, five are from the mountains both to the north and to the south of the
isolated Cerro Piedra Larga, (Chiropterotriton sp., Abronia oaxacae, Sceloporus formosus,
Eumeces brevirostris, and Thamnophis godmani). Five are known only from the
mountains to the north (Hyla bistincta, Sceloporus grammicus, Anolis quercorum,
Rhadinaea fulvivittis, and Crotalus ravus), whereas the other three are known only
from the mountains to the south. Hence, the montane herpetofauna of Cerro Piedra
Larga shows affinities to both mountain systems. For this reason we can not say
Cerro Piedra Larga represents a characteristic local fauna. It is worth mentioning
that the Chiropterotriton may represent an undescribed species and that in all
likelihood the populations from the mountains north of Cerro Piedra Larga are
specifically distinct from those to the south.
The herpetofauna of the dry forest shows strong influence of the Isthmus of
Tehuantepec. Indeed,of the 21 species found in that habitat, 13 are also found on
the Isthmus (Hartweg & Oliver 1940). Most likely, the dry valley of the Río
Tehuantepec acts as a dispersal corridor for these faunas, whereas the long and
complex series of habitat types connecting the area to the Oaxaca Valley may serve
as an effective filter barrier to dispersal.
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
451
In total, five species endemic to the state of Oaxaca were detected in the area:
three in dry forest, and two in pine-oak forest. For example, Abronia oaxacae is
restricted to the pine-oak forests of the Sierra Madre del Sur (Bogert & Porter
1967). Ctenosaura oaxacana is a recently described species for the Isthmus of
Tehuantepec, but was long referred to C. quinquecarinata by many herpetologists
(Bailey 1928, Gicca 1983, de Quieroz 1995).
Ecotones between two habitats can hold both mixtures of the faunas of each, or
can possess distinct faunal elements (Cox & Moore 1980, Pianka 1982). In the
ecotonal zone on Cerro Piedra Larga, we encountered only four species, three of
which were also found in dry forest (Sceloporus siniferus, Lepidophyma smithi and
Leptotyphylops goudoti). One species, the turtle Rhynoclemys rubida, was found only in
the ecotone, although the species has been found at a nearby locality (El Camarón)
in dry forest, suggesting that the species would also be found in that habitat near
Cerro Piedra Larga.
Altitudinal distributions of species tend to follow particular patterns in Mexico
(Wake & Lynch 1976, Flores V. 1993, Peterson et al. 1993), such as that species
richness declines with elevation. This pattern was present, although not marked,
in the Cerro Piedra Larga herpetofauna. Twenty-one species were found in the
low-elevation dry forest, whereas 15 species were found in the high-elevation pineoak forest. Because inventory intensity was greatest in the pine-oak forest, we suspect
that this pattern would be even more pronounced if the inventories were completed.
The diversity of microhabitats present in an area, or spatial heterogeneity, is an
important factor in determining how many species may live in an area (Cox &
Moore 1980). Within the broad category of ground surface microhabitats, for
example, the lizard Sceloporus siniferus was found in seven distinct microhabitats,
which is probably why this species is found across a broad swath of elevations.
Others are much more specific, such as the plethodontid salamander Chiropterotriton
sp., which was found only in bromeliads, as is usual for the group (Wake 1987).
Birds. A total of 84 species was encountered in the humid montane forests on Cerro
Piedra Larga (Appendix 3). Of these, 59 probably breed on the mountain, and the
remaining 25 are passage migrants or winter residents. Faunal accumulation curves
(Fig. 3) leveled off after 11 days, indicating that the inventory effort was more or
less complete.
By far the dominant species, both in mist nets and in observations, was the
nightingale-thrush Catharus occidentalis; other common species included the
hummingbird Hylocharis leucotis, the jay Cyanocitta stelleri, the warbler Basileuterus
belli, and the brush-finch Atlapetes brunneinucha. The list for the site also includes
several rare and poorly known species (Binford 1989), such as the Maroon-fronted
Ground-dove Claravis mondetoura, the Military Macaw Ara militaris, and the Northern
Saw-whet owl Aegolius acadicus.
Cerro Piedra Larga, therefore, represents the last island of humid montane
forest before the lowland barrier of the Isthmus of Tehuantepec. Cerro Piedra
452
A. TOWNSEND PETERSON ET AL.
Larga, at 95 48' W longitude, lies isolated between two mountain ranges, one to
the north and one to the south (Fig. 1). Hence, this island is both marginal and
isolated between the two coastal mountain ranges, and Cerro Piedra Larga records
of many montane forest species represent considerable range extensions south
and east towards the Isthmus of Tehuantepec (e.g., Cyrtonyx montezumae, Dendrortyx
macroura, Otus flammeolus, Aegolius acadicus, Catharus occidentalis, and Ridgwayia
pinicola, among others).
Cerro Piedra Larga’s avifauna includes most of the species typical of humid
montane forests in southern Mexico (Binford 1989, Navarro-Sigüenza 1992, TorresChávez 1992, Hernández-Baños et al. 1995). Species occurring in similar habitats
on other mountain ranges west of the Isthmus of Tehuantepec in Oaxaca, but not
detected on Cerro Piedra Larga, include Otus trichopsis, Glaucidium gnoma, Strix
varia, Amazilia beryllina, Eupherusa spp., Tilmatura dupontii, Atthis heloisa, Anabacerthia
variegaticeps, Automolus rubiginosus, Xiphocolaptes promeropirhynchus, Xiphorhynchus
flavigaster, Empidonax affinis, E. difficilis, E. fulvifrons, Cyanolyca spp., Catharus
aurantiirostris, Dendroica graciae, Myioborus miniatus, Piranga flava, P. erythrocephala,
Chlorospingus ophthalmicus, Icterus graduacauda, Loxia curvirostra, and Carduelis notata.
Of these species, several are rare and often difficult to detect (e.g. Tilmatura dupontii,
Strix varia) or show seasonal movements (e.g. Amazilia beryllina, Carduelis notata);
others perhaps require the existence of humid forests at adjacent lower elevations
(e.g. Anabacerthia variegaticeps, Eupherusa spp.). However, our failure to detect several
species almost certainly reflects their absence from the humid montane forests of
Cerro Piedra Larga: Xiphorhynchus flavigaster, Myioborus miniatus, and most notably
Chlorospingus ophthalmicus, all of which are common and easily detectable when
present in an area.
Two peculiar features of the avifauna of Cerro Piedra Larga deserve further
comment. First, very few representatives of the neotropical families
Dendrocolaptidae (only Lepidocolaptes affinis) and Furnariidae (none) were found;
even the neotropical flycatchers (Tyrannidae) seemed less common than on other
mountain ranges in Oaxaca. This imbalance has also been noted on other islands
of humid montane forest isolated amid dry forest (Morales-Pérez & NavarroSigüenza 1991).
Second, the relative abundances of the nightingale-thrushes Catharus occidentalis
and C. frantzii were strikingly uneven. We netted C. occidentalis 69 times, and C.
frantzii but twice, a 35-fold difference. These two species were for many years
confused by systematists, and often occur syntopically (Phillips 1969), but their
abundances seem rarely to be balanced. We have detected no obvious feature of
vegetation or habitat type associated with which species is dominant, so further
study of the habitat use and ecological requirements of this species pair will be
necessary to understand factors affecting their relative abundances.
The position of Cerro Piedra Larga intermediate between the mountains of
eastern and western Mexico was the reason that we chose the mountain for careful
study. The arid pine-oak forest connection to the eastern end of the Sierra de
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
453
Miahuatlán (Fig. 1), led us to expect affinities with the Sierra Madre del Sur of
southern Oaxaca and Guerrero. However, our study of the specimens resulting
from the inventory did not support this prediction.
For seven species, it was possible to determine the subspecific affinities and
distinguish between eastern and western origins of the Cerro Piedra Larga
populations. The quail Cyrtonyx montezumae is known in Oaxaca only from the Sierra
Madre del Sur and the mountains of the interior (Binford 1989), and hence is of
western affinities. The quail-dove Claravis mondetoura, although not previously
known from Oaxaca, has been recorded in Mexico only in Chiapas and southern
Veracruz (Howell 1992), and hence has decidedly eastern affinities. The owl Aegolius
acadicus is known in Oaxaca reliably only from the vicinity of Cerro San Felipe in
the northern interior of the state (Binford 1989), a forest island belonging to the
interior-western group identified by Hernández-Baños et al. (1995). For the
hummingbird species complex Lampornis amethystinus, which is represented by forms
with pink or blue throats distributed in a mosaic across southern Mexico (Phillips
1966), we collected examples of both forms from the same net line, providing the
first evidence of sympatry of the two forms (Torres-Chávez et al. in prep.), and
indicating the presence of populations of western (L. [a.] margaritae) and of largely
eastern (L. [a.] amethystinus) affinity. The toucanet Aulacorhynchus [prasinus] prasinus
of the east, separable at the species level from the western Mexican populations A.
[p.] wagleri (Navarro-Sigüenza et al. 2002), was identified on the basis of remains in
the posession of local residents. Finally, the antpitta Grallaria [guatimalensis]
ochraceiventris of the west, separable at the species level from the eastern G. [g.]
guatimalensis (Navarro-Sigüenza et al. in prep.) was collected. Therefore, in all, we
encountered four forms with affinities in the west or interior of Oaxaca, and three
forms with affinities to eastern Oaxaca, and the avifauna of Cerro Piedra Larga is
seen to consist of a mixture of the two avifaunas.
The discovery and documentation of the avifauna of Cerro Piedra Larga as a
mixture of eastern and western avifaunas changes considerably the picture of
isolated insular (montane) faunas (Hernández-Baños et al. 1995). Formerly, the
two faunas were almost without exception thought to be allopatric, and any isolated
peak would be quickly determinable as “eastern” or “western,” on the basis of a few
records. Cerro Piedra Larga, however, combines elements of both faunas, probably
owing to random colonization from mountains to the north and south across the
dry, lowland barriers. For the first time, then, montane avifaunas in Mexico are
seen to show mobility and ability to colonize remote regions, and form novel faunistic
combinations.
Mammals. We recorded 15 species of four orders, nine families and 16 genera
(Appendix 4). Bats were the most diverse group, with eight species, followed by
rodents (four species), artiodactyls (two species), and insectivores (one species).
Both localities sampled showed a mixture of pure pine-oak forest with restricted
patches of cloud forest. In all, ten species were of nearctic distribution, being
454
A. TOWNSEND PETERSON ET AL.
distributed principally to the north of Cerro Piedra Larga (e.g., Cryptotis mexicana,
Neotoma mexicana, Pteronotus parnelli, Mormops megalophylla), whereas three
(Dermanura azteca, Carollia perpicilliata, Sturnira lilium) are of neotropical distribution
(Goodwin 1969). Overall, the mammal fauna at Cerro Piedra Larga showed a
similar pattern of mixture of nearctic and neotropical affinities, as in other mountain ranges (Briones et al. 2001, Sánchez-Cordero 2001).
Discussion and conclusions
We present herein a preliminary biotic inventory of Cerro Piedra Larga, an isolated massif in Oaxaca, southern Mexico. This massif, which had not previously
seen any detailed floristic or faunistic study, resulted quite interesting for a number of reasons. Concerted study of several taxonomic groups made the results all
the more intriguing.
The first surprise was the existence of patches of cloud forest at high elevations
on such an isolated interior mountain massif. This habitat, which includes a number
of floristic elements characteristic of cloud forests elsewhere, is otherwise restricted
in Oaxaca to the Sierra Madre Oriental, the Sierra Madre del Sur, and a few isolated
mountains in northern Oaxaca (Rzedowski 1978, Binford 1989). Hence, the
discovery of cloud forest patches on Cerro Piedra Larga provides a fascinating
isolated example of this habitat.
Perhaps most interesting was the pattern of geographic affinities of the highelevation faunas on Cerro Piedra Larga. Among birds, reptiles, and amphibians,
we were able to distinguish a series of species that have clear affinities to eastern
Mexico (northern Oaxaca) or western Mexico (southern Oaxaca). The usual
understanding of cloud forest biogeography emphasizes vicariance, with areas
relatively continuous in the Pleistocene being subdivided under warmer climate
regimes during interglacial periods. This scenario would predict previous
connection of mountain masses over the past several tens of thousands of years,
and would suggest sharing of faunas with the regions with which connections were
most recently shared (Hernández-Baños et al. 1995).
The pattern of affinities observed on Cerro Piedra Larga, however, did not fit
this pattern. The quite-distinct faunas of the Sierra Madre Oriental versus the Sierra
Madre del Sur each have contributed species to the fauna of this massif, and the
mountain’s faunas are seen to be a mixture of the two faunas. This result indicates
a new process in montane vertebrate biogeography: colonization across dry lowland
barriers.
Acknowledgments. This study was made possible by the generosity of our field companions,
particularly Tom Gnoske. Funding was provided by the U.S. National Science Foundation.
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
455
Literature cited
BAILEY, J. W. 1928. A revision of the lizard genus Ctenosaura. Proceedings of the U.S. National
Museum 73: 1-55.
BINFORD, L. C. 1989. A distributional survey of the birds of the Mexican state of Oaxaca.
Ornithological Monographs 43: 1-405.
BOGERT, C. M. & A. P. PORTER. 1967. A new species of Abronia (Sauria, Anguidae) from the
Sierra Madre del Sur of Oaxaca. American Museum Novitates 2279: 1-29.
BRIONES, M., V. SÁNCHEZ-CORDERO & G. QUINTERO. 2001. Listado de los mamíferos terrestres
del norte del estado de Oaxaca. Anales del Instituto de Biologia, Universidad Nacional
Autónoma de México, Serie Zoología 72: 125-161.
CAMBELL, J. A. & D. R. FROST. 1993. Anguid lizards of the genus Abronia: Revisionary notes,
description of four new species, a phylogenetic analysis, and a key. Bulletin of the American
Museum of Natural History 216: 1-121.
CASAS-ANDREU, G., F. R. MÉNDEZ DE LA CRUZ & J. L. CAMARILLO-RANGEL. 1996. Anfíbios y
reptiles de Oaxaca: lista, distribución y conservación. Acta Zoológica Mexicana 69: 1-35.
COX, C. B. & P. D. MOORE. 1980. Biogeography: an ecological and evolutionary approach, 3rd ed.
Blackwell Scientific Publications, London.
DARDA, D. M. 1994. Alloenzyme variation and morphological evolution among Mexican
salamanders of the genus Chiropterotriton (Caudata: Plethodontidae). Herpetologica 50:
164-187.
DE QUIEROZ, K. 1995. Checklist and key to the species of Mexican iguanas (Reptiles:
Iguanidae). Publicaciones Especiales del Museo de Zoología, Facultad de Ciencias 9: 1-48.
DUELLMAN, W. E. 1966. The Central American herpetofauna: an ecological perspective.
Copeia 1966: 700-719.
DUELLMAN, W. E. 2001. Hylid frogs of Middle America. Society for the Study of Amphibians
and Reptiles, Ithaca, New York.
FLORES V., O. A. 1993. Herpetofauna of Mexico: distribution and endemism. In: T. P.
Ramamoorthy, R. Bye, A. Lot & J. Fa (eds.) Biological diversity of Mexico: origins and
distribution. Oxford University Press, Oxford, pp. 253-279.
FLORES V., O. A. & A. MUÑOZ. 1993. Anfíbios y reptiles. In: Luna V. I. & J. E. Llorente B.
(eds.) Historia Natural del Parque Ecológico Estatal Omiltemi, Chilpancingo, Guerrero, México.
UNAM/CONABIO, México, D. F.
GICCA, D. F. 1983. Enyaliosaurus quinquecarinatus. Catalog of American amphibians and reptiles
329: 1-2.
GOODWIN, G. G. 1969. Mammals from the state of Oaxaca, Mexico, in the American Museum
of Natural History. Bulletin of the American Museum of Natural History 141: 1-269.
HARTWEG, N. & J. A. OLIVER. 1940. A contribution to the herpetology of the Isthmus of
Tehuantepec, IV. Miscellaneous Publications of the Museum of Zoology, University of Michigan
47: 1-33.
HERNÁNDEZ M., J. C. 1992. Herpetofauna del Municipio de San Cristóbal de las Casas. San Cristóbal,
Chiapas.
HERNÁNDEZ-BAÑOS, B. E., A. T. PETERSON, A. G. NAVARRO-SIGÜENZA & P. ESCALANTE-PLIEGO.
1995. Bird faunas of the humid montane forests of Mesoamerica: biogeographic patterns and conservation priorities. Bird Conservation International 5: 251-277.
456
A. TOWNSEND PETERSON ET AL.
HOWELL, S. N. G. 1992. Recent records of Maroon-chested Ground-Dove in Mexico. Euphonia
1: 39-41.
MITTERMEIER, R. A., N. MYERS, J. B. THOMSEN & G. A. B. D. FONSECA. 1998. Biodiversity
hotspots and major tropical wilderness areas: approaches to setting conservation priorities. Conservation Biology 12: 516-520.
MORALES-PÉREZ, J. E. & A. G. NAVARRO-SIGÜENZA. 1991. Análisis de distribución de las aves
en la Sierra Norte del estado de Guerrero, México. Anales del Instituto de Biologia,
Universidad Nacional Autónoma de México, Serie Zoología 62: 497-510.
NAVARRO-SIGÜENZA, A. G. 1992. Altitudinal distribution of birds in the Sierra Madre del Sur,
Guerrero, México. Condor 94: 29-39.
NAVARRO-SIGÜENZA, A. G., A. T. PETERSON, E. LÓPEZ-MEDRANO & H. BENÍTEZ-DÍAZ. 2002.
Species limits in Mesoamerican Aulacorhynchus toucanets. Wilson Bulletin 113: 363-372.
PETERSON, A. T., O. A. FLORES V., L. S. LEÓN P., J. E. LLORENTE B., M. A. LUIS M., A. G.
NAVARRO-SIGÜENZA, M. G. TORRES CH. & I. VARGAS F. 1993. Conservation priorities in
northern Middle America: moving up in the world. Biodiversity Letters 1: 33-38.
PETERSON, A. T., A. G. NAVARRO-SIGÜENZA & H. BENÍTEZ-DÍAZ. 1998. The need for continued
scientific collecting: a geographic analysis of Mexican bird specimens. Ibis 140: 288-294.
PHILLIPS, A. R. 1966. Further systematic notes on Mexican birds. Bulletin of the British Ornithologists’ Club 86: 86-94.
PHILLIPS, A. R. 1969. An ornithological comedy of errors: Catharus occidentalis and C. frantzii.
Auk 86: 605-623.
PIANKA, E. R. 1982. Evolutionary ecology. Harper & Row, London.
PISANI, G. R. & J. VILLA. 1974. Guía de técnicas de preservación de anfíbios y reptiles. SSAR
Misc. Publ. Circ. Herp 2: 1-24.
RAMAMOORTHY, T. P., R. BYE, A. LOT & J. FA (eds.). 1993. Biological diversity of Mexico: origins
and distribution. Oxford University Press, Oxford.
RZEDOWSKI, J. 1978. Vegetación de México. Limusa, México, D.F.
SÁNCHEZ, O. & W. LÓPEZ-FORMENT. 1988. Anfíbios y reptiles de la región de Acapulco,
Guerrero, México. Anales del Instituto de Biologia, Universidad Nacional Autónoma de México,
Serie Zoología 58: 735-750.
SÁNCHEZ-CORDERO, V. 2001. Elevational gradients for bats and rodents. Global Ecology and
Biogeography 10: 63-76.
TORRES-CHÁVEZ, M. G. 1992. Distribución altitudinal de las aves en la Sierra de Juárez, Oaxaca.
Universidad Nacional Autónoma de México, México, D. F.
VOGT, R. G. & R. L. HINE. 1982. Evaluation of techniques for assessment of amphibian and
reptile populations in Wisconsin. In: N. J. J. Scott (ed.) Herpetological communities. U. S.
Department of the Interior, Fish and Wildlife Service, Washington, D.C., pp. 201-217
WAKE, D. B. 1987. Adaptive radiation of salamanders in Middle American cloud forest.
Annals of the Missouri Botanical Garden 74: 242-264.
WAKE, D. B. & J. F. LYNCH. 1976. The distribution, ecology, and evolutionary history of
plethodontid salamanders in tropical America. Bulletin of the Los Angeles County Museum
of Natural History 25: 1-65.
WILSON, L. D. 1990. Tantilla striata. Catalog of American amphibians and reptiles 504.
Recibido: 1.IX.2003
Aceptado: 10.II.2004
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PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
Appendix 1. List of plant species encountered on Cerro Piedra Larga, Oaxaca, Mexico.
Species
Actinidaceae
Saurauia scabrida Hemsl.
Amaranthaceae
Iresine celosia L.
Alstroemeriaceae
Bomarea hirtella (Kunth) Herb.
Aquifoliaceae
Ilex discolor Hemsl.
I. quercetorum I.M. Johnst.
Araliaceae
Oreopanax xalapensis (Kunth) Decne. et Planch.
Asclepiadaceae
Gonolobus sp.
Marsdenia macrophylla (Humb. et Bonpl. ex Schult.)
E. Fourn.
Sarcostemma sp.
Asteraceae
Calea scabra (Lag.) B.L. Rob.
Erigeron karvinskianus DC.
Eupatorium chiapense B.L. Rob.
Eupatorium cremastum B.L. Rob.
Eupatorium mairetianum DC.
Eupatorium pycnocephalum Less.
Eupatorium pazcuarense Kunth
Eupatorium sp.
Gnaphalium americanum Mill.
Gnaphalium attenuatum DC.
Gnaphalium roseum Kunth
Hieracium irazuense Benth.
Hieracium sp.
Mikania pyramidata Donn. Sm.
Rumfordia attenuata B.L. Rob.
Schistocarpha seleri Rydb.
Senecio aspatensis (Coult.) Greenm.
Senecio jurgensenii Hemsl.
Senecio polypodioides Greene
Stevia triflora DC.
Verbesina nelsonii B. L. Rob. et Greenm.
Begoniaceae
Begonia fusca Liebm.
BPQS
BPQH
BMM
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
BPQS = arid pine-oak forest, BPQH = humid pine-oak forest, and BMM = cloud
forest.
458
A. TOWNSEND PETERSON ET AL.
Appendix 1. Continues.
Species
Betulaceae
Alnus acuminata Kunth subsp. arguta (Schltdl.) Furlow
Alnus jorullensis Kunth
Bromeliaceae
Catopsis sp.
Pitcairnia sp.
Buddlejaceae
Buddleia crotonoides A. Gray
Cactaceae
Aporocactus flagelliformis (L.) Lem.
Caprifoliaceae
Viburnum discolor Benth.
Caryophyllaceae
Arenaria lanuginosa (Michx.) Rohrb.
Stellaria cuspidata Willd. ex Schltdl.
Chloranthaceae
Hedyosmum mexicanum C. Cordem.
Clethraceae
Clethra licanioides Standl. et Steyerm.
Clethra macrophylla M. Martens et Galeotti
Cornaceae
Cornus disciflora DC.
Cornus excelsa Kunth
Crassulaceae
Sedum sp.
Cucurbitaceae
Cyclanthera ribiflora (Schltdl.) Cogn.
Cistaceae
Helianthemum pringlei S. Watson
Dryopteridaceae
Dryopteris pseudofilix-mas (Fée) Rothm.
Phanerophlebia macrosora (Baker) Underw.
Polystichum distans E. Fourn.
Ericaceae
Arbutus glandulosa M. Martens et Galeotti
Arctostaphylos aff. longifolia Benth.
A. pyrifolia (Donn. Sm. ex Loes.) Standl. et Steyerm.
Gaultheria odorata Bredem. ex Willd.
Vaccinium confertum Kunth
Xolisma squamulosa (M. Martens et Galeotti) Small
Fagaceae
Quercus candicans Née
Q. crassifolia Humb. et Bonpl.
BPQS
X
BPQH
BMM
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
459
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
Appendix 1. Continues.
Species
BPQS
Q. laurina Bonpl.
Q. scytophylla Liebm.
Q. uxoris McVaugh
Garryaceae
Garrya laurifolia Hartw. ex Benth.
Gesneriaceae
Moussonia deppeana (Schltdl. & Cham.) Hanst.
Iridaceae
Orthrosanthus monadelphus Ravenna
Lamiaceae
Salvia gracilis Benth.
S. lavanduloides Kunth
S. mexicana L.
S. aff. recurva Benth.
Satureja macrostema (Benth.) Briq.
Stachys repens M. Martens et Galeotti
Lauraceae
Litsea glaucescens Kunth
Leguminosae
Astragalus guatemalensis var. brevidentatus (Hemsl.) Barneby
Crotalaria mollicula Kunth
Inga sp.
Lotus repens (G. Don.) Sessé et Moc. ex Standl. et Steyerm.
Lupinus sp.
Lentibulariaceae
Pinguicula moranensis Kunth
Lobeliaceae
Lobelia gruina Cav.
L. laxiflora Kunth
Melastomataceae
Miconia glaberrima (Schltdl.) Naudin
M. mexicana (Bonpl.) Naudin
Tibouchina galeottiana (Triana) Cogn.
Myricaceae
Myrica cerifera L.
Myrsinaceae
Ardisia nigropunctata Oerst.
Rapanea juergensenii Mez
Myrtaceae
Eugenia oerstediana O. Berg
Nolinaceae
Nolina longifolia (Karw. ex Schult. f.) Hemsl.
X
Onagraceae
Fuchsia arborescens Sims
BPQH
BMM
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
460
A. TOWNSEND PETERSON ET AL.
Appendix 1. Continues.
Species
Fuchsia sp.
Lopezia racemosa Cav.
Orchidaceae
Isochilus unilateralis B.L. Rob.
Lemboglossum maculatum (La Llave et Lex.) Halb.
Lemboglossum cervantesii (La Llave et Lex.) Halb.
Passifloraceae
Passiflora membranacea Benth.
P. sexflora Juss.
Pinaceae
Pinus ayacahuite C. Ehrenb. ex Schltdl.
P. douglasiana Martínez
P. lawsonii Roezl ex Gordon
P. maximinoi H. E. Moore
P. oocarpa Schiede ex Schltdl.
Piperaceae
Peperomia sp.
Piper amalago L.
P. scabrum Sw.
Polygalaceae
Monnina xalapensis Kunth
Ranunculaceae
Anemone mexicana Kunth
Rosaceae
Holodiscus argenteus (L.f.) Maxim.
Prunus brachybotrya Zucc.
P. salasii Standl.
Rubiaceae
Galium mexicanum Kunth
Hoffmannia conzattii B.L. Rob.
Nertera granadensis (Mutis ex L. f.) Druce
Palicourea padifolia (Willd. ex Roem. et Schult.)
C.M. Taylor et Lorence
Psychotria pubescens Sw.
Relbunium hypocarpium (L.) Hemsl.
Saxifragaceae
Heuchera sp.
Scrophulariaceae
Calceolaria mexicana Benth.
Smilacaceae
Smilax regelii Killip et C.V. Morton
Solanaceae
Cestrum anagyris Dunal
Cestrum sp.
BPQS
BPQH
X
X
X
X
BMM
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
461
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
Appendix 1. Continues.
Species
BPQS
Lycianthes amatitlanensis (J.M. Coult. et Donn. Sm.) Bitter
Physalis amphitricha (Bitter) Standl. et Steyerm.
P. philadelphica Lam.
Solandra sp.
Solanum aligerum Schltdl.
X
S. appendiculatum Kunth ex Dunal
S. lanceolatum Cav.
S. pubigerum Dunal
S. nigricans M. Martens et Galeotti
S. tacanense Lundell
Styracaceae
Styrax argenteus C. Presl.
S. polyneurus Perkins
Theaceae
Cleyera theaeoides (Sw.) Choisy
Urticaceae
Myriocarpa longipes Liebm.
Valerianaceae
Valeriana palmeri A. Gray
Violaceae
Viola guatemalensis W. Becker
BPQH
X
X
X
X
X
BMM
X
X
X
X
X
X
X
X
X
X
X
X
X
X
462
A. TOWNSEND PETERSON ET AL.
Appendix 2. Distribution of species of amphibians and reptiles by vegetation type,
altitudinal interval, and use of microhabitat types on Cerro Piedra Larga, Oaxaca.
Species
Bufonidae
Bufo canaliferus
B. marinus
Hylidae
Hyla bistincta
H. sumichrasti
H. sp.
Ptychohyla leonhardschultzei
Leptodactylidae
Eleutherodactylus pipilans
Plethodontidae
Chiropterotriton sp.
Anguidae
Abronia oaxacae
Eublepharidae
Coleonyx elegans
Gekkonidae
Phyllodactylus muralis
Iguanidae
Anolis quercorum
Ctenosaura oaxacana
Sceloporus formosus
S. grammicus
S. siniferus
S. smithi
Urosaurus bicarinatus
Scincidae
Eumeces brevirostris
Teiidae
Aspidoscelis deppii
A. guttata
A. motaguae
Xantusiidae
Lepidophyma smithi
Colubridae
Senticolis triapis
Enulius flavitorques
Leptophis diplotropis
Rhadinaea fulvivittis
Microhabitat
Pineoak
Ff
Ff
Ff
x
x
x
Ff
Dry
forest
Ecotone
Elevation
(m)
x
x
670-1005
725
x
x
x
2425
850
1375-1670
1375-1670
x
1020-1030
A
x
2600
A
x
2405-2580
A
x
850
Ff
x
1005
Ff, A
A
Ff, A
A
Ff
Ff
Ff
x
Ff
x
x
x
x
x
x
x
1140-1855
Ff
Ff
Ff
x
x
x
Ff
x
Ff
A
Ff
Ff= Forest floor; A= Arboreal; F= fossorial
x
x
x
x
1850-2095
725-980
2200-2540
2285-2610
950-1765
725-1000
840-945
x
x
830-1005
920-1000
1005
x
1350-1590
2425
950
1605-2145
2405-2580
463
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
Appendix 2. Continues.
Species
Tantilla striata
Thamnophis godmani
Trimorphodon biscutatus
Leptotyphlopidae
Leptotyphlops goudoti
Elapidae
Micrurus ephippifer
Viperidae
Crotalus ravus
Bataguridae
Rhynoclemmys rubida
Microhabitat
Ff
Ff
Pineoak
x
Ff
Dry
forest
Ecotone
x
1000
2580-2610
845
x
x
Elevation
(m)
x
1005-1225
Ff
x
1610
Ff
x
1590
Ff
x
1351
464
A. TOWNSEND PETERSON ET AL.
Appendix 3. Bird species recorded in humid montane forests on Cerro Piedra Larga, Oaxaca,
31 March - 13 April 1993. Species likely to breed on the mountain are indicated in the
“Breeds” column.
Species
Accipiter striatus
Aegolius acadicus
Aeronautes saxatalis
Ara militaris
Atlapetes brunneinucha
A. pileatus
Aulacorhynchus prasinus
Basileuterus belli
Bombycilla cedrorum
Buteo brachyurus
B. jamaicensis
Buteogallus anthracinus
Caprimulgus vociferus
Cardellina rubrifrons
Cathartes aura
Catharus frantzii
C. guttatus
C. occidentalis
Catherpes mexicanus
Certhia americana
Chaetura vauxi
Claravis mondetoura
Colaptes [auratus] cafer
Colibri thalassinus
Columba fasciata
Contopus pertinax
C. sordidulus
Coragyps atratus
Corvus corax
Cyanocitta stelleri
Cyrtonyx montezumae
Dendroica auduboni
D. occidentalis
D. townsendi
Dendrortyx macroura
Diglossa baritula
Empidonax sp.
Breeds Mar 31 Apr 1 2
?
Y
Y
Y
Y
Y
Y
Y
N
Y
Y
Y
?
N
Y
Y
N
Y
Y
Y
?
?
Y
Y
Y
Y
N
Y
Y
Y
Y
N
N
N
Y
Y
?
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
0
0
0
1
0
1
1
0
1
0
1
1
0
1
0
0
1
1
1
1
1
0
1
1
1
0
1
1
0
0
1
0
1
1
1
1
1
0
1
0
1
1
1
1
0
1
0
1
1
0
1
0
1
1
1
0
0
1
0
1
1
1
1
0
0
3
4
5
6
7
8 9 10 11 12 13
0
0
1
0
1
1
1
1
1
0
1
0
1
0
1
1
0
1
0
1
1
0
1
0
1
1
0
0
0
1
0
1
1
1
1
0
0
0
0
0
0
1
1
1
1
0
1
1
1
1
0
1
1
1
1
0
1
1
0
1
0
1
1
0
1
1
1
0
0
1
1
1
1
1
0
1
0
1
1
1
1
1
1
0
1
0
1
1
1
1
1
1
0
1
1
0
1
0
1
0
0
0
0
1
1
1
1
1
0
0
1
0
0
1
1
1
1
0
1
1
0
0
0
1
0
1
0
1
1
0
1
0
0
0
0
1
0
0
0
0
1
1
0
1
1
0
0
1
1
0
0
1
1
1
0
1
0
0
1
0
1
0
0
0
0
1
0
1
0
0
0
1
1
0
0
1
0
1
1
0
1
1
0
0
0
0
0
0
1
1
1
1
1
0
0
0
0
1
1
1
0
0
1
0
1
0
1
0
0
1
0
0
1
0
1
1
0
1
1
0
0
0
0
0
0
1
1
1
0
1
1
0
1
0
1
0
0
0
1
1
0
1
0
0
1
1
1
0
0
0
0
1
1
1
1
1
1
0
0
0
0
0
1
1
1
0
1
1
0
0
0
1
0
1
0
1
1
1
1
0
1
0
0
1
0
1
0
0
1
1
1
1
1
1
0
1
0
0
1
1
1
1
0
0
1
1
1
0
0
0
1
0
0
1
0
1
0
1
0
0
1
0
0
1
0
1
1
1
1
1
0
0
1
0
0
1
1
1
1
0
0
1
0
1
0
0
0
1
0
1
1
0
1
0
0
0
1
1
0
0
0
0
1
1
1
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
0
0
0
0
0
0
0
465
PRELIMINARY BIOLOGICAL SURVEY, CERRO PIEDRA LARGA OAXACA
Appendix 3. Continues.
Species
Ergaticus ruber
Eugenes fulgens
Geococcyx velox
Grallaria guatimalensis
Hirundo pyrrhonota
H. rustica
Hylocharis leucotis
Icterus [galbula] bullockii
Junco phaeonotus
Lampornis amethystinus
L. margaritae
Lepidocolaptes affinis
Leptotila verreauxi
Melanerpes formicivorus
Melanotis caerulescens
Micrastur semitorquatus
Mitrephanes phaeocercus
Myadestes occidentalis
Myiarchus tuberculifer
Myioborus pictus
Oporornis tolmiei
Otus flammeolus
Parula superciliosa
Peucedramus taeniatus
Pheucticus ludovicianus
P. melanocephalus
Picoides villosus
Pipilo ocai
Piranga ludoviciana
Ptilogonys cinereus
Regulus calendula
Ridgwayia pinicola
Sphyrapicus varius
Stelgidopteryx serripennis
Streptoprocne zonaris
Tachycineta thalassina
Troglodytes brunneicollis
Trogon mexicanus
Breeds Mar 31 Apr 1 2
Y
Y
Y
Y
N
N
Y
N
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
N
Y
Y
Y
N
Y
Y
Y
N
Y
N
Y
N
?
Y
N
Y
Y
0
0
0
0
0
0
1
0
1
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
1
1
1
0
0
1
0
0
1
0
0
1
0
1
0
0
1
1
0
0
0
0
0
0
1
0
1
1
0
0
1
1
1
0
0
0
0
0
1
0
1
0
0
1
0
1
1
0
1
1
1
0
0
1
1
0
0
0
1
0
1
1
1
0
1
1
0
1
0
1
3
4
5
6
7
8 9 10 11 12 13
1
0
0
1
0
0
1
0
1
0
0
1
0
1
1
1
1
1
0
0
0
1
1
0
0
0
1
1
0
1
1
0
0
0
0
0
1
1
1
1
0
0
0
0
1
0
1
0
0
1
0
1
1
0
1
1
0
0
0
0
1
0
0
0
1
1
0
1
1
0
0
0
0
0
0
1
1
0
0
0
0
0
1
0
0
0
0
1
0
1
1
0
1
1
0
0
0
0
1
0
0
0
1
0
0
1
1
0
1
0
0
1
1
1
1
0
0
0
0
0
1
0
0
0
0
1
0
1
1
0
0
1
0
0
0
0
1
0
0
1
1
1
1
0
1
0
0
0
0
0
0
1
1
1
0
0
0
0
1
0
0
1
0
1
0
0
1
0
0
1
0
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
0
0
1
1
1
0
0
0
0
0
1
0
0
1
0
1
0
0
1
0
1
1
0
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
1
1
1
1
1
0
0
0
0
0
1
0
0
1
0
1
0
0
1
0
1
1
0
0
1
0
1
0
0
0
1
0
1
1
1
0
1
0
0
0
1
1
1
0
1
0
1
1
1
0
0
1
0
1
1
0
1
0
1
1
0
0
1
0
1
0
0
1
1
0
1
1
1
1
0
0
1
1
1
1
1
1
0
0
0
0
1
0
0
1
1
1
0
0
1
0
1
1
1
0
1
0
1
0
0
1
1
0
0
1
1
1
1
0
1
0
1
1
1
0
0
0
0
0
1
0
0
1
0
1
0
1
1
0
1
1
0
0
0
0
1
1
1
1
1
1
1
1
1
1
0
0
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
466
A. TOWNSEND PETERSON ET AL.
Appendix 3. Continues.
Species
Turdus infuscatus
T. migratorius
Vermivora celata
V. ruficapilla
Vireo huttoni
V. solitarius
Wilsonia pusilla
Zonotrichia lincolnii
Breeds Mar 31 Apr 1 2
Y
Y
N
N
Y
N
N
N
0
1
0
0
0
0
0
0
1
1
0
0
0
1
1
1
1
1
1
0
0
1
1
1
3
4
5
6
7
8 9 10 11 12 13
1
1
0
0
0
0
1
0
1
1
1
0
1
0
1
0
1
1
0
0
0
0
1
0
1
1
0
0
0
0
1
0
1
1
0
0
0
0
1
0
1
1
0
0
1
1
1
0
1
1
0
1
1
0
1
0
1
1
0
0
1
0
1
0
1
1
0
0
1
0
1
0
1
1
0
0
1
1
1
0
1
0
0
0
0
0
0
0
Appendix 4. List of mammal species encountered on Cerro Piedra Larga, Oaxaca, México.
ORDER INSECTIVORA
Cryptotis mexicana (Coues, 1877)
ORDER CHIROPTERA
Pteronotus parnelli (Gray, 1843)
Glossophaga soricina (Pallas, 1766)
Dermanura azteca (K. Andersen, 1906)
Carollia perspicillata (Linnaeus, 1758)
Sturnira lilium (E. Geoffroy, 1810)
Tadarida brasilensis (I. Geoffroy, 1824)
Mormops megalophylla (Peters, 1864)
Myotis vellifer (Kaup, 1929)
ORDER RODENTIA
Reithrodontomys fulvescens (J.A. Allen, 1894)
Oryzomys alfaroi (J.A. Allen, 1891)
Neotoma mexicana (Baird, 1855)
Sciurus aureogaster (F. Cuvier, 1829)
ORDER ARTIODACTYLA
Odocoileus virginianus (Zimmermann, 1815)
Pecari tayacu (Linnaeus, 1758)